Solanum evolvulifolium, Greenm.,

Tepe, EJ & Bohs, L, 2011, A revision of Solanum section Herpystichum, Systematic Botany 36 (4), pp. 1068-1087 : 1077-1079

4. SOLANUM EVOLVULIFOLIUM Greenm.,

Bot. Gaz. 37: 211. 1904.

—TYPE: COSTA RICA. San José: La Palma , 1,460 m, Sep 1898 (fl, fr), A. Tonduz 12615 (lectotype, here designated: GH–GH00077489 !; possible isolectotypes: CR–12615, K–K000449423!, K–K000449424!, M–M0111204!, NY–NY00138984!, US – US 00027570!).

Vine, sometimes shrubby, climbing tree trunks or other vegetation. Stems slender, woody, glabrous to densely pubescent, the trichomes typically ca. 1 mm or longer, occasionally in distinct lines along stem. Sympodial units plurifoliate. Leaves simple, the arrangement distinctly distichous, the blades 0.5–5 × 0.3–3 cm, 1–3 times as long as wide, ovate to oblong-ovate, chartaceous to coriaceous, sand-punctate, glabrous to densely pubescent on the leaf blade adaxially and abaxially, pubescent on midvein adaxially with hairs 0.2–0.5 mm, discolored, occasionally reddish below; venation pinnate, with 3–4 pairs of secondary veins, the veins densely sand-punctate; base truncate to cordate, often oblique; margins entire to undulate, revolute; apex rounded and apiculate to acute, sometimes acuminate; petioles nearly absent to 0.5 cm, glabrous to densely pubescent, densely sand-punctate. Internodes 0.5– 1.5(–3.5) cm. Inflorescences 1–15 cmlong, unbranched, nearly leaf-opposed to extra-axillary, with 2–80 flowers (scars), the axes glabrous to pubescent with simple, uniseriate, curled hairs; peduncle 0.6–1.5 cm; rachis 1–14 cm; pedicels 4– 10 mm in flower, green to pink, 10–15 mm in fruit, glabrous to densely pubescent, spaced nearly contiguously to 8 mm apart. Calyx 1.5–3 mm long, conical, the tube 1–2.5 mm long, the lobes 0.5–1.5 × 0.8–1.5 mm, deltate, rounded and minutely apiculate at tips to acute, the margins somewhat thickened, glabrous to sparsely pubescent along margins, more dense at tips of the lobes, pale green to pink; fruiting calyx minutely accrescent. Corolla 1–2 cmindiameter, 5–8 mm long, stellate, membranous, greenish, white, pink, to bluish-purple, sometimes mottled, the lobes 5–11 × 2–3 mm, lanceolate, acute at apices, glabrous adaxially and abaxially, the margins ciliate.Stamens withfilamentsca. 1 mm long, glabrous; anthers 2–3 × 0.8–1.5 mm. Ovary glabrous; style 4–6 × ca. 0.25 mm, cylindrical to somewhat clavate, minutely papillose in lower 2/3; stigma truncate to somewhat capitate. Fruit 0.6–1.5 × 0.6–1 cm, globose to ovoid, slightly flattened, rounded to acute at apex, glabrous, red to reddish-brown when ripe. Seeds 1.5–2 mm in diameter, rounded, tan, the surface rugulose. Figure 1H–I .

Habitat and Distribution— Solanum evolvulifolium occursin Costa Rica, Panama, Colombia, Venezuela, Ecuador, and Peru as an epiphyte on tree trunks in rain and cloud forest habitats; (200–) 800–2,600 m in elevation ( Fig. 6 ).

Phenology— Flowering and fruiting apparently occurs yearround.

Etymology— The epithet evolvulifolium refers to the similarity of the leaves of this species to some species of Evolvulus (Convolvulaceae).

Notes— Solanum evolvulifolium , a climbing species, is recognizable by its characteristic distichous leaf arrangement ( Fig. 1H ) and branching pattern. The main stem of this species is most often encountered climbing on tree trunks, attached with adventitious roots at the nodes; secondary branches extend away from the main stem. Higher order branches are typically distichous and arise at ca. 45° angles, often giving the plant a characteristic, flattened appearance. The leaves often diminish in size along a branch, occasionally to a branching point or inflorescence, and then increase in size again. The pedicels, calyx and corolla are frequently pinkish-white to greenish-pink. This species can be distinguished from other simple leaved, viny members of sect. Herpystichum by the uniformly short internodes, somewhat coriaceous leaves (instead of chartaceous or somewhat fleshy), and the flattened aspect of the branches that results from the distichous leaves branches.

South American collections are more variable than those from Central America in leaf shape and, especially, the degree of pubescence. The calyx lobes of Central American collections are broadly ovate and rounded apically, whereas the lobes of most South American collections are deltoid in shape and acute apically. These calyx characters, however, are not sufficiently uniform, nor are they correlated with other characters that might justify segregation of S. evolvulifolium into one or more additional species. Several extreme forms from South America are included here, and these include especially robust forms from Colombia and Ecuador and an especially pubescent form from Ecuador. It is possible that the differences merit specific status, but both forms are represented by only a few collections and are thus included within a broadly defined and variable, yet easily identifiable S. evolvulifolium . Furthermore, the three sequenced accessions of S. evolvulifolium , including one accession of the robust form and two of the standard form, form a monophyletic group together with S. crassinervium and S. loxophyllum ( Fig. 4 ). The robust form of S. evolvulifolium is strongly supported as sister to Central American accessions of S. evolvulifolium lending support for its inclusion in a morphologically variable, yet easily recognizeable S. evolvulifolium .

There is some confusion about the numbering and collector of the lectotype collection of S. evolvulifolium . Greenman’s protologue states that the collector and number is Pittier 7413. However, the handwritten labels on the K specimens give the collector as Tonduz. They do not have a collector number, but they do have a “herb. nat. Cost. number 12615 (Herbario del Museo Nacional de Costa Rica). Nevertheless, there is sufficient overlap of label information on different specimens to determine that they are all clearly part of the same collection. All labels list Tonduz’ name and the herb. nat. Cost. number. Furthermore, it seems that Tonduz was the actual collector, but that Pittier distributed the specimens under his own set of numbers ( Dauphin López 2009; B. Hammel, pers. comm.). For these reasons, we refer to the collection as Tonduz 12615.

Greenman cited two syntypes, Tonduz 12615 (as Pittier 7413) and Wercklé 11599, in his original description of S. evolvulifolium . Tonduz 12615 is chosen as the lectotype because this collection has many duplicates, whereas only a single specimen is known of the Wercklé collection. The GH specimen of Tonduz 12615 is chosen as the lectotype because Greenman was at GH in 1904 and this is likely the specimen that he used in his description of S. evolvulifolium . Furthermore, the GH specimen gives the altitude as 1460 m, the altitude stated in the protologue. The rest of the Tonduz 12615 collections report an altitude of 1,542 m and therefore they are listed above as possible isolectotypes.

Representative Specimens Examined— COSTA RICA. La Palma, 1,500 m, Nov 1897 (fr), K. Wercklé 11599 ( GH). Alajuela: Reserva Biológica Monteverde, 10°19’N 84°43’W, 820 m, 22 Oct 1988 (fl), E. Bello 468 (F, MO); Reserva Forestal de Arenal, 10°18’N 84°42’W, 850–900 m, 28 Feb 1990 (fl), E. Bello 1961 ( MO); San Carlos, La Forma, Finca El Jilguero, 10°25′25″N 84°42′05″W, 800 m, 22 Nov 1992 (fl), G. Herrera 5696 ( MO); 12 km NNW of San Ramón by road on way to San Lorenzo, 1 km Sof Balsa, 10°10’N 84°29’W, 1,100 m, 25 Apr 1983 (fl), R. L. Liesner & E. Judziewicz 14936 (MO, WIS). Cartago: Beside Río Villegas, valley of Río Grande de Orosi, 9°42’N 83°47’W, 1,620 m, 11 Jan 1970 (fl), R. W. Lent 1849 (F, MO, U); Monumento Nacional Guayabo, 9°58’N 83°41’W, 26 Jan 1993 (fl), G. Rivera 2054 (F, K); growing in dense upland rain forest about 5 km SW of Tapanti, 9°47’N 83°55’W, 1,500 m, 17 Aug 1967 (fr), J. Taylor & C. Taylor 4472 (MO, NY). Heredia: Vicinity of Colonia Virgen del Socorro, 10°17’N 84°10’W, 900 m, 10 Aug 1975, (fl), J. Utley & K. Utley 2824 (F); Parque Nacional Braulio Carrillo, 10°15′50″N 84°05’W, 1,200 –1,400 m, 13 Nov 1986 (fl), M. Grayum & G. Herrera 7881 ( MO). Limon: Guápiles, Los Angeles, San Miguel, 10°04′20″N 83°50′40″W, 1,300 m, 21 Feb 1990 (fl), A. Chacón et al. 744 ( MO); Parque Nacional Cordillera de Talamanca, 9°22′30″N 83°14′10″W, 1,700 m, 24 Mar 1993 (fl), A. Fernández 818 (BM, MO); El Progreso, 9°47′20″N, 83°07′30″W, 1,600 m, 24 Apr 1989 (fl), G. Herrera & A. Chacón 2771 (F, MO, NY); San Jose: road between Alto La Palma and Bajo La Hondura, 1,400 m, 24 Feb 1978 (fl), F. Almeda & K Nakai 3913 ( MO); Pérez Zeledón, Savegre, 9°31’N 83°51’W, 1,900 m, 3 Aug 1994 (fl, fr), G. Herrera et al.7253 (K); woods near Quebrada Vargus, Alto La Palma, 10°04’N 83°59’W, 1,400 m, 31 Mar 1974 (fl), R. W. Lent 3852 (AAU, F, MO).

PANAMA. Bocas del Toro: Nslope of Cerro Horqueta, 8°49′24”N 82°26′54″W, 6,000 –7,000 ft, 5 Aug 1947 (fr), P. H. Allen 4995 (F, G, U); along road from Fortuna Dam, towards Chiriquí Grande, 2.2 miles Nof the continental divide, 8°45’N 82°15’W, 800 m, 12 Mar 1985 (fl), G. McPherson 6815 (MO, WIS). Chiriqui: Boquete, Bajo Chorro, 8°50’N 82°29’W, 6,000 ft, 12 Jan 1938 (fl) M. E. Davidson 112 (GH, F, US). Chiriquí/Bocas del Toro: Zona Protectora Palo Seco, along continental divide, 8°47’N 82°13’W, 1,100 –1,300 m, 11 Aug 2000 (fl), S. Knapp & J. Mallet 9178 (BR, MO).

COLOMBIA. Antioquia: Río Caldera, 5°58’N, 74°58’W, Jan 1953 (fl), Bro. Daniel 4498 ( US). Caldas: La Finca, Quindío, 4°27’N 75°40’W, 3,200 m, Feb 1937 (st), E. Dryander 2136 ( US); Santa Cecelia, Cordillera occidental,vertiente occidental, 5°18’N 76°13’W, 800 m, 29 Dec 1945 (fl), K. von Sneidern 5524 (F). Cauca: El Tambo, Parque Nacional Munchique, 2°36′40”N 74°54′10″W, 2,570 m, 20 Jul 1993 (fl, fr), G. Lozano et al. 6961 (COL, MA); El Cairo, Cerro del Inglés, Cordillera Occidental, 4°45’N 76°13’W, 2,260 m, 5 Jan 1987 (fl), F. A. Silverstone Sopkin et al. 2967 ( NY); El Tambo, La Costa, 4°4’N 77°1’W, 1,500 m, 25 Mar 1938 (infl), K.von Sneidern 1663 (F, G, S, US). Chocó: San Jose del Palmar, Hoya del Rio Torito, Finca Los Guaduales, 5°02’N 76°22’W, 630 m, 6 Mar 1980 (fl), E. Forero et al. 6793 (COL, MO); 6 km Eof Río Pato, ca. 48 km Wof Las Animas on the Pan American Highway, 5°20N 76°49’W, 250 m, 11 Jan 1979 (fl, fr), A. H.Gentry & A. Renteria 24016 (AAU, MO). Nariño: Cordillera Occidental, Finca La Planada, near Chucunes, 1°11’N 77°58’W, 1,950 m, 13 Jan 1981 (fr), A. H. Gentry et al. 30549 (COL, MO, NY); Junin-Barbacoas road, 2–10 km. Nof Junin, 1°30’N 78°10’W, 900 m, 26 Jul 1986, (st), A. H. Gentry et al. 55333 ( MO). Putumayo: Villagarzón, Carretera a Pto. Asis, 4 May 1994 (st), J. L. Fernández 11431 ( COL); Punto de Buenos Aires, Cerro Portachuelo, 0°22’N 75°01’W, 2,080 m, 27 Jul 1964 (fl), D. D. Soejarto 1139 ( GH). Risaralda: Apía, Vereda La Cumbre, 5°8’42N 76°0′46″W, 2,285 m, 24 Feb 1983 (fl), J.H.Torres 2216 ( COL); Pereira, La Pastora, 2,500 m, 20 Jan 1998 (fl), G. Vargas 4472 ( COL). Valle: Cordillera occidental, Hoya del Río Digua, La Elsa, 3°15’N 70°51’W, 1,000 –1,200 m, 9 Nov 1943 (fl, fr), J. Cuatrecasas 15313 (F, US).

VENEZUELA. Mérida: Rivas Dávila, 22–27 km Sof Tovar along rd. to Canaguá, 8°14’N 71°45’W, 2,100 –2,256 m, 16 Apr 1984 (fl, fr), J. L. Luteyn et al. 9960 ( NY).

ECUADOR. Carchi: Mira, Norte del Carmen, Camino a Chical, 0°17’N 78°13’W, 2,000 m, 10 Feb 1992 (fl), W. A. Palacios et al. 9753 ( MO); Tulcan, Reserva Indígena Awá, 0°53’N 78°25’W, 1,800 m, 17 Aug 1992 (fr), G. Tipaz et al. 1923 ( MO). Carchi / Esmeraldas: Near Lita, 0°52’N 78°27’W, 600 m, 19 May 1987 (fl). H. Van der Werff et al. 9493 ( MO). Cotopaxi: Sigchos, Triunfo Grande, 0°32′22”S 78°58′59″W, 2,349 m, 6 Aug 2003 (fl, fr), J. E. Ramos Perez et al.7061 ( NY); Pujilí,Reserva Biológica Los Ilinizas, 0°58′45”S 79°06′53″W, 1,725 m, 10 Aug 2003 (st), F. A. Silverstone Sopkin et al. 10027 (MO, NY). Esmeraldas: Road Lita-Alto Tambo, ca. km 17.8, 0°51’N 78°29’W, 850 m, 28 Sep 1991 (fl), B. Ollgaard 99154 ( AAU). Loja: Cerro de Celica, Celica - Guachanamá, km 2.7, 4°05′46”S 79°56′45″W, 2,250 m, 12 Apr 1994 (fl, fr), P. M. Jorgensen et al. 93 ( NY). Napo: Parque Nacional Sumaco y Comunidad de Pacto Sumaco, 0°38′56″S 77°35′49″W, 1,550 –1,700 m, 26 Apr 1997 (fr), A. Alvarez et al. 2017 ( MO); Quijos, Cosanga, Yanayacu Biological Station and Center for Creative Studies, 0°35′55”S 77°53′22″W, 2,200 m, 18 Aug 2005 (fl), J. L. Clark et al. 9438 (BM, NY, US); Km 2, carretera nueva Cotundo – Coca, 0°42’N 77°42’W, 1,130 m, 5 Aug 1984 (fl), C. H. Dodson et al. 15031 (NY, MO).Pichincha:Reservaecológica Río Guajalito, km 59 delacarretera antigua Quito-St. Domingo de los Colorados, 0°13′53″S 74°48′10″W, 1,800 – 2,000 m, 4 May 2000 (fl, fr), I. Tapia 1254 ( SEL). Sucumbios: El Reventador, colecciones en ámbas márgenes del Rio Reventador, 0°02’N 77°41’W, 1,850 m, 6 Oct 1990 (st), J. Jaramillo & E. Grijalva 12933 (AAU, NY).

PERU. Amazonas: Monte Virgen, 50 m frente la comunidad de Caterpiza, 3°55’S 77°42’W, 200 m, 30 Aug 1979, (fl, fr), V. Huashikat 268 ( MO).

Figures

FIG.1. Flowers fruits, and habit of Solanum sect. Herpystichum: A. Solanum limoncochaense, habit, flower, and bud. B. S. limoncochaense fruit in situ and C. in cross section (E. J. Tepe & S. Stern 2627, QCNE). D. S. trifolium flower and leaf (E. J. Tepe & S. Stern 2682, QCNE). E. S. phaseoloides habit, flower, and leaf (photo by W. H. Haber). F. S. crassinervium habit (E. J. Tepe & S. Stern 2729, QCNE), and G. flowers (S. Stern & E. J. Tepe 400, QCNE). H. S. evolvulifolium habit and flower, and I. fruit (immature) (E. J. Tepe & S. Stern 2671, QCNE). J. S. pacificum inflorescence with flowers, buds, and fruit (immature), and K. habit (E. J. Tepe et al. 2696, QCNE). L. S. loxophyllum habit, note adventitious roots at nodes, and M. older stem with inflorescences, note figure eightshaped stem (E. J. Tepe & S. Stern 2726, QCNE). Inflorescences emerge from the cleft between the two lobes of the stem.

FIG.6. Distribution of Solanum evolvulifolium.

FIG.4. Summary phylogeny of Solanum sect. Herpystichum based on based on plastid psbA-trnH, trnT-trnF, and trnS-trnG, and nuclear waxy, ITS, cos1C, cos5, cos9B, cos10B, and cos11 sequences (E. J. Tepe et al. 2011). For clarity, replicate accessions of all species except for S. evolvulifolium and S. pentaphyllum have been omitted. Provenance is indicated for species with multiple accessions, species with compound leaves are underlined, and the asterisk indicates an especially robust specimen of S. evolvulifolium. The topology is based on Bayesian analyses, and branch support values are Bayesian posterior probabilities/maximum parsimony bootstrap.